Genomic regions influencing gene expression of the HMW glutenins in wheat

Bread wheat (Triticum aestivum L.) produces glutenin storage proteins in the endosperm. The HMW glutenins confer distinct viscoelastic properties to bread dough. The genetics of HMW glutenin proteins have been extensively studied, and information has accumulated about individual subunits, chromosomal locations and DNA sequences, but little is known about the regulators of the HMW glutenins. This investigation addressed the question of glutenin regulators. Expression of the glutenins was analyzed using QRT-PCR in ditelosomic (dt) Chinese Spring (CS) lines. Primers were designed for each of 4 CS glutenin genes and a control, non-storage protein endosperm-specific gene Agp-L (ADP-glucose pyrophosphorylase). Each line represents CS wheat, lacking one chromosome arm. The effect of a missing arm could feasibly cause an increase, decrease or no change in expression. For each HMW glutenin, results indicated there were, on average, 8 chromosome arms with an up-regulatory effect and only one instance of a down-regulatory effect. There were significant correlations between orthologous and paralogous HMW glutenins for effects of chromosome groups B and D. Some or all the glutenin alleles shared regulatory loci on chromosome arms 2BS, 7BS, 4DS, 5DS and 6DS, and Agp-L shared regulatory loci with glutenins on arms 7AS, 7BS, 2DS, 3DS, 4DS and 5DS. These results suggest a few chromosome arms contain putative regulatory genes affecting the expression of conserved cis elements of 4 HMW glutenin and Agp-L genes in CS. Regulation by common genes implies the regulators have diverged little from the common wheat ancestor, and furthermore, some regulation may be shared by endosperm-specific-genes. Significant common regulators have practical implications.     

Chromosomal locations of the genes for histones and a histone gene-binding protein family HBP-1 in common wheat

The chromosomal locations of the genes in common wheat that encode the five histones and five members of the HBP (histone gene-binding protein)-1 family were determined by hybridizing their cloned DNAs to genomic DNAs of nullitetrasomic and telosomic lines of common wheat, Triticum aestivum cv. Chinese Spring. The H1 and H2a genes are located on different sets of homoeologous chromosomes or chromosome arms, namely, 5A, 5B and 5D, and 2AS, 2BS and 2DS, respectively. Genes for the other histones, H2b, H3 and H4, are found in high copy number and are dispersed among a large number of chromosomes. The genes for all members of the HBP-1 family are present in small copy numbers. Those for HBP-1a(1) are located on six chromosome arms, 3BL, 5AL, 5DL, 6AL, 6BS and 7DL, whereas those for each HBP-1a(c14), 1a(17), 1b(c1), and 1b(c38) are on a single set of homoeologous chromosome arms; 4AS, 4BL, 4DL; 6AS, 6BS, 6DS; 3AL, 3BL, 3DL; and 3AS, 3BS, 3DS, respectively. The genes for histones H1 and H2a, and for all members of the HBP-1 family except HBP-1a(1) are assumed to have different phylogenetic origins. The genes for histone 2a and HBP-1a(17) are located in the RFLP maps of chromosomes 2B and 6A, respectively. Gene symbols are proposed for all genes whose chromosomal locations have been determined.     

Genetic characterization of QTL associated with resistance to Fusarium head blight in a doubled-haploid spring wheat population.

Fusarium head blight (FHB) is one of the most important fungal wheat diseases worldwide. Understanding the genetics of FHB resistance is the key to facilitating the introgression of different FHB resistance genes into adapted wheat. The objectives of the present study were to detect and map quantitative trait loci (QTL) associated with FHB resistance genes and characterize the genetic components of the QTL in a doubled-haploid (DH) spring wheat population using both single-locus and two-locus analysis. A mapping population, consisting of 174 DH lines from the cross between DH181 (resistant) and AC Foremost (susceptible), was evaluated for type I resistance to initial infection during a 2-year period in spray-inoculated field trials, for Type II resistance to fungal spread within the spike in 3 greenhouse experiments using single-floret inoculation, and for resistance to kernel infection in a 2001 field trial. One-locus QTL analysis revealed 7 QTL for type I resistance on chromosome arms 2DS, 3AS, 3BS, 3BC (centromeric), 4DL, 5AS, and 6BS, 4 QTL for type II resistance on chromosomes 2DS, 3BS, 6BS, and 7BL, and 6 QTL for resistance to kernel infection on chromosomes 1DL, 2DS, 3BS, 3BC, 4DL, and 6BS. Two-locus QTL analysis detected 8 QTL with main effects and 4 additive by additive epistatic interactions for FHB resistance and identified novel FHB resistance genes for the first time on chromosomes 1DL, 4AL, and 4DL. Neither significant QTL by environment interactions nor epistatic QTL by environment interactions were found for either type I or type II resistance. The additive effects of QTL explained most of the phenotypic variance for FHB resistance. Marker-assisted selection for the favored alleles at multiple genomic regions appears to be a promising tool to accelerate the introgression and pyramiding of different FHB resistance genes into adapted wheat genetic backgrounds.     

Chromosomal locations of genes that control major RNA-degrading activities in common wheat (Triticum aestivum L.)

Seventeen RNA-degrading enzymes of common wheat, with apparent molecular masses from 42.2 kDa to 16.3 kDa, were observed by the RNA-SDS-PAGE assay. To determine their chromosome locations, all chromosome arms of common wheat except 4BS were assayed in their null condition by using a set of ditelosomic or nullitetrasomic lines of the cultivar Chinese Spring. Our results showed that only one chromosome location each was identified for the 22.8-kDa and the 21.2-kDa enzymes, as well as for the 21.6 kDa enzyme, and they are on chromosome arms 2AS and 2DS, respectively. Loci controlling the 20.1 kDa activity were on chromosome arms 2AL, 4BS, 4DS and 6BS. The locus or loci coding for the gene(s) of the 42.2-kDa, 40.9-kDa and 39.2-kDa enzymes were probably ocated on chromosome arm 5AS, and their expression, in agreement with most other RNA-degrading enzyme activities were stimulated when chromosome arm 5AL was missing, indicating a inhibiting locus on 5AL. Our data suggested that the 31.9-kDa, 30.6-kDa and 29.6-kDa enzymes were possibly products of a common precursor which might be coded by a gene(s) on chromosome arm 6BS, and that the processing is co-regulated by loci on chromosome arms 2BS, 3DS, 6AL, 6BL and 7BS. The remaining of the enzyme activities were consistently found in all of the lines tested, and thus are presumably encoded by multiple loci. The only other possibility is that, their loci may be on chromosome arm 4BS which we have not assayed in its null condition.Contribution from Agriculture Research Division, University of Nebraska. Journal Series No. 11271 Current address: Dept. Bio/Microbiology, South Dakota State University, Brookings, SD57007, USA     

A cytogenetic ladder-map of the wheat homoeologous group-4 chromosomes

We report the results of chromosome maps of wheat homoeologous chromosomes 4A, 4B, and 4D using 40 RFLP markers and 39 homozygous deletion lines. Deletion breakpoints divide the chromosomes into 45 subarm intervals with 32 intervals distinguished by molecular markers. The chromosome maps confirm the homoeology of arms 4AS to 4BL and 4DL, and 4AL to 4BS and 4DS. The chromosome map of 4A reveals novel information concerning the 4AL-5AL-7BS cyclical translocation. The presence of homoeologous group-4 long-arm markers, Xksu G10 and Xpsr 1051, intervening between the translocated 5AL and 7BS chromosome segments in 4AL suggests that the translocation events are more complex than was earlier believed. Chromosome maps confirm a pericentric inversion in Chinese Spring chromosome 4B. The consensus chromosome map is compared to the genetic map of wheat to construct a cytogenetic ladder-map (CLM). The CLM reveals an unequal distribution of recombination along the length of the chromosome arms. Recombination is highest in the distal half, and low in the proximal half, of the chromosome arms.     

Molecular Mapping of Wheat: Major Genes and Rearrangements in Homoeologous Groups 4, 5, and 7

A molecular-marker linkage map of hexaploid wheat (Triticum aestivum L. em. Thell) provides a framework for integration with the classical genetic map and a record of the chromosomal rearrangements involved in the evolution of this crop species. We have constructed restriction fragment length polymorphism (RFLP) maps of the A-, B-, and D-genome chromosomes of homoeologous groups 4, 5, and 7 of wheat using 114 F(7) lines from a synthetic X cultivated wheat cross and clones from 10 DNA libraries. Chromosomal breakpoints for known ancestral reciprocal translocations involving these chromosomes and for a known pericentric inversion on chromosome 4A were localized by linkage and aneuploid analysis. Known genes mapped include the major vernalization genes Vrn1 and Vrn3 on chromosome arms 5AL and 5DL, the red-coleoptile gene Rc1 on 7AS, and presumptively the leaf-rust (Puccinia recondita f.sp. tritici) resistance gene Lr34 on 7DS and the kernel-hardness gene Ha on 5DS. RFLP markers previously obtained for powdery-mildew (Blumeria graminis f.sp. tritici) resistance genes Pm2 and Pm1 were localized on chromosome arms 5DS and 7AL.     

小麦新种质241千粒重的基因定位

以小麦新种质材料241为测定系,中国春单体系及双端体系作为测验系,对控制千粒重的基因进行了定位研究,结果表明,小麦新种质材料241的3A,2B,7B和6D染色体上具有控制千粒重的基因,其中3A,7B和6D染色体上的基因表现为强效, 2B染色体上的基因表现为弱效,通过双端体分析进一步将控制千粒重的基因定位到3AL,7BL和6DL上,其中6DL上可能具有控制241千粒重的新基因。     

DNA marker analysis for pyramided of Fusarium head blight (FHB) resistance QTLs from different germplasm

A pyramided FHB resistance line of wheat (WSY) was previously developed from three FHB resistant cultivars (Sumai 3, Wangshuibai, and Nobeokabouzu) in the Jiangsu Academy of Agricultural Sciences, China. In the present study, we analyzed the genetic relationship between WSY and the three parental cultivars using DNA markers in order to clarify how many and which resistance genes had accumulated in WSY. We analyzed 282 DNA markers from the 21 wheat chromosomes. WSY was found to include different chromosome regions that harbored putative FHB QTLs of the three parental germplasm. Haplotypes of DNA markers on these QTL regions revealed that the 1BL, 2BL, 5AS, and 7AL QTL regions were from Sumai 3, the 2AS, 2DS, 3AS, and 6BS QTL regions were from Wangshuibai, and the 3BS QTL region was from Nobeokabouzu. This study showed that different resistance genes from the different resistant germplasm had indeed accumulated in WSY. WSY is a potential resistant resource for FHB resistance in wheat breeding programs.     

Seed-storage-protein loci in RFLP maps of diploid, tetraploid, and hexaploid wheat

 Linkages between high- and low-molecular-weight (M_r) glutenin, gliadin and triticin loci in diploid, tetraploid and hexaploid wheats were studied by hybridization of restriction fragments with DNA clones and by SDS-PAGE. In tetraploid and hexaploid wheat, DNA fragments hybridizing with a low-M_r glutenin clone were mapped at the XGlu-3 locus in the distal region of the maps of chromosome arms 1AS, 1BS, and 1DS. A second locus, designated XGlu-B2, was detected in the middle of the map of chromosome arm 1BS completely linked to the XGli-B3 gliadin locus. The restriction fragments mapped at this locus were shown to co-segregate with B subunits of low-M_r glutenins in SDS-PAGE in tetraploid wheat, indicating that XGlu-B2 is an active low-M_r glutenin locus. A new locus hybridizing with the low-M_r clone was mapped on the long arm of chromosome 7A^m in diploid wheat. No glutenin protein was found to co-segregate with this new locus. Triticin loci were mapped on chromosome arms 1AS, 1BS, and 1DS. A failure to detect triticin proteins co-segregating with DNA fragments mapped at XTri-B1 locus suggests that this locus is not active. No evidence was found for the existence of Gli-A4, and it is concluded that this locus is probably synonymous with Gli-A3. Recombination was observed within the multigene gliadin family mapped at XGli-A11 (1.2 cM).1 Although these closely linked loci may correspond to the previously named Gli-A1 and Gli-A5 loci, they were temporarily designated XGli-A1.1 and XGli-A1.2 until orthology with Gli-A1 and Gli-A5 is established. Received: 25 March 1997 / Accepted: 23 June 1997     

Chromosomal control of glutenin subunits in aneuploid lines of wheat: analysis by reversed-phase high-performance liquid chromatography

Summary Glutenin subunits from nullisomic-tetrasomic and ditelocentric lines of the hexaploid wheat variety ‘Chinese Spring’ (CS) and from substitution lines of the durum wheat variety ‘Langdon’ were fractionated by reversed-phase high-performance liquid chromatography (RP-HPLC) at 70 °C using a gradient of acetonitrile in the presence of 0.1% trifluoroacetic acid. Nineteen subunits were detected in CS. The presence and amounts of four early-eluted subunits were found, through aneuploid analysis, to be controlled by the long arms of chromosomes 1D (1DL) (peaks 1–2) and 1B (1BL) (peaks 3–4). Sodium dodecyl sulfate-polyacrylamide gel electrophoresis showed that these four subunits are the high molecular weight subunits of glutenin, which elute in the order 1Dy, 1Dx, 1By, and 1Bx. Similar amounts of 1DL subunits were present (6.3 and 8.8% of total glutenin), but 1BL subunits differed more in abundance (5.4 and 9.5%, respectively). Results indicate that most late-eluting CS glutenin subunits were coded by structural genes on the short arms of homoeologous group 1 chromosomes: 6 by 1DS, 5 by 1AS, and 4 by 1BS. Glutenin of tetraploid ‘Langdon’ durum wheat separated into nine major subunits: 6 were coded by genes on 1B chromosomes, and 3 on 1A chromosomes. Gene locations for glutenin subunits in the tetraploid durum varieties ‘Edmore’ and ‘Kharkovskaya-5’ are also given. These results should make RP-HPLC a powerful tool for qualitative and quantitative genetic studies of wheat glutenin. The mention of firm names or trade products does not imply that they are endorsed or recommended by the U.S. Department of Agriculture over other firms or similar products not mentioned Stationed at the Northern Regional Research Center, Peoria.     

Linkage mapping of genes controlling endosperm storage proteins in wheat

Summary A translocation mapping procedure was used to map gene-centromere distances for the genes controlling endosperm proteins on the short arm of each of the chromosomes 1A, 1B and 1D in wheat. The genes controlling triplet proteins (tentatively designated Tri-1) were found to be closely linked to the centromere on chromosome arms 1AS and 1DS and loosely linked to the gliadin genes (Gli-1) on the same arms. The Gli-1 genes segregated independently or were very loosely linked to their respective centromeres. The Gli-B1-centromere map distance on 1BS was also estimated using conventional telocentric mapping and the result was similar to that obtained with the translocation mapping. A simple two-step one-dimensional electrophoretic procedure is described which allows the low-molecular-weight (LMW) glutenin subunits to be separated from the gliadin bands, thus facilitating the genetic analysis of these LMW subunits. No recombination was observed between the genes (designated Glu-3) controlling some major LMW glutenin subunits and those controlling gliadins on chromosome arms 1AS and 1DS. However, in a separate experiment, the genes controlling LMW glutenin subunits on 1BS (Glu-B3) showed a low frequency of recombination with the gliadin genes.Portion of the Ph.D. thesis submitted by the senior author     

Microsatellites confirm the authenticity of inter-varietal chromosome substitution lines of wheat (Triticum aestivum L.)

Ninety-five wheat microsatellite markers (WMS) were used to verify the authenticity of the set of Saratovskaya 29/Yanetzkis Probat inter-varietal wheat chromosome substitution lines developed using Saratovskaya 29 as the recipient variety. Polymorphic markers were available for all chromosome arms except 4DS, 6DS and 7DS. Each chromosome substitution line was tested by 2–8 microsatellite markers. The results demonstrate that most of the lines are correct. Out of 21 lines tested 17 showed the expected microsatellite pattern of the donor variety. Two entire chromosomes, 1B and 7A, and two chromosome arms, 3AL and 6DS, were not substituted with Yanetzkis Probat in their respective lines. Three microsatellite markers located in the distal regions of chromosome arms 4AL, 3BS and 5BL in the corresponding substitution lines did not reveal the expected microsatellite pattern of the recipient variety. The possible causes of the incorrect substitution line development and the appearance of incorrect distal microsatellite markers are discussed. The data confirm the idea that microsatellite markers provide ideal tools for testing the authenticity of genetic stocks of wheat. Received: 27 August 1999 / Accepted: 8 October 1999     

Quantitative Trait Loci for Aluminum Resistance in Wheat Cultivar Chinese Spring

Aluminum (Al) toxicity is one of the major constrains for wheat production in many wheat growing areas worldwide. Further understanding of inheritance of Al resistance may facilitate improvement of Al resistance of wheat cultivars (Triticum aestivum L.). A set of ditelosomic lines derived from the moderately Al-resistant wheat cultivar Chinese Spring was assessed for Al resistance. The root growth of ditelosomic lines DT5AL, DT7AL, DT2DS and DT4DS was significantly lower than that of euploid Chinese Spring under Al stress, suggesting that Al-resistance genes might exist on the missing chromosome arms of 5AS, 7AS, 2DL and 4DL of Chinese Spring. A population of recombinant inbred lines (RILs) from the cross Annong 8455 × Chinese Spring-Sumai 3 7A substitution line was used to determine the effects of these chromosome arms on Al resistance. A genetic linkage map consisting of 381 amplified fragment length polymorphism (AFLP) markers and 168 simple sequence repeat (SSR) markers was constructed to determine the genetic effect of the quantitative trait loci (QTLs) for Al resistance in Chinese Spring. Three QTLs, Qalt.pser-4D, Qalt.pser-5A and Qalt.pser-2D, were identified that enhanced root growth under Al stress, suggesting that inheritance of Al resistance in Chinese Spring is polygenic. The QTL with the largest effect was flanked by the markers of Xcfd23 and Xwmc331 on chromosome 4DL and most probably is multi-allelic to the major QTL identified in Atlas 66. Two additional QTLs, Qalt.pser-5A and Qalt.pser-2D on chromosome 5AS and 2DL, respectively, were also detected with marginal significance in the population. Some SSR markers identified in this study would be useful for marker-assisted pyramiding of different QTLs for Al resistance in wheat cultivars.     

Mapping of QTLs affecting copper tolerance and the Cu,Fe, Mn and Zn contents in the shoots of wheat seedlings

Quantitative trait loci (QTLs) for Cu-tolerance were determined in wheat grown in control and Cu-treated soil in greenhouse. In addition, loci having an influence on the shoot Cu-, Fe-, Mn-and Zn-contents under non-stressed and Cu-stressed environments were mapped. One major QTL for Cu-tolerance was found on chromosome 5DL, while slighter effects were determined on the chromosomes 1AL, 2DS, 4AL, 5BL and 7DS. QTLs affecting the shoot Mn-and Zn-contents were found on the chromosomes 3BL and 3AL, respectively. The centromeric region on the chromosome 3B plays a role in the regulation of the shoot Fe-contents in the stressed plants. Under Cu-stress QTL affecting shoot Cu-content was found on chromosome 1BL, while on the chromosome 5AL a QTL influencing the Cu-accumulation ability of wheat from Cu-polluted soil was determined.     

Genome-wide association mapping for resistance to leaf rust,stripe rust and tan spot in wheat reveals potential candidate genes

Key message

Genome-wide association mapping in conjunction with population sequencing map and Ensembl plants was used to identify markers/candidate genes linked to leaf rust, stripe rust and tan spot resistance in wheat.

Abstract

Leaf rust (LR), stripe rust (YR) and tan spot (TS) are some of the important foliar diseases in wheat (Triticum aestivum L.). To identify candidate resistance genes for these diseases in CIMMYT’s (International Maize and Wheat Improvement Center) International bread wheat screening nurseries, we used genome-wide association studies (GWAS) in conjunction with information from the population sequencing map and Ensembl plants. Wheat entries were genotyped using genotyping-by-sequencing and phenotyped in replicated trials. Using a mixed linear model, we observed that seedling resistance to LR was associated with 12 markers on chromosomes 1DS, 2AS, 2BL, 3B, 4AL, 6AS and 6AL, and seedling resistance to TS was associated with 14 markers on chromosomes 1AS, 2AL, 2BL, 3AS, 3AL, 3B, 6AS and 6AL. Seedling and adult plant resistance (APR) to YR were associated with several markers at the distal end of chromosome 2AS. In addition, YR APR was also associated with markers on chromosomes 2DL, 3B and 7DS. The potential candidate genes for these diseases included several resistance genes, receptor-like serine/threonine-protein kinases and defense-related enzymes. However, extensive LD in wheat that decays at about 5?×?10⁷ bps, poses a huge challenge for delineating candidate gene intervals and candidates should be further mapped, functionally characterized and validated. We also explored a segment on chromosome 2AS associated with multiple disease resistance and identified seventeen disease resistance linked genes. We conclude that identifying candidate genes linked to significant markers in GWAS is feasible in wheat, thus creating opportunities for accelerating molecular breeding.

     

Identification of novel QTLs for seedling and adult plant leaf rust resistance in a wheat doubled haploid population

Pyramiding of genes that confer partial resistance is a method for developing wheat (Triticum aestivum L.) cultivars with durable resistance to leaf rust caused by Puccinia triticina. In this research, a doubled haploid population derived from the cross between the synthetic hexaploid wheat (SHW) (×Aegilotriticum spp.) line TA4152-60 and the North Dakota breeding line ND495 was used for identifying genes conferring partial resistance to leaf rust in both the adult plant and seedling stages. Five QTLs located on chromosome arms 3AL, 3BL, 4DL, 5BL and 6BL were associated with adult plant resistance with the latter four representing novel leaf rust resistance QTLs. Resistance effects of the 4DL QTL were contributed by ND495 and the effects of the other QTLs were contributed by the SHW line. The QTL on chromosome arm 3AL had large effects and also conferred seedling resistance to leaf rust races MJBJ, TDBG and MFPS. The other major QTL, which was on chromosome arm 3BL, conferred seedling resistance to race MFPS and was involved in a significant interaction with a locus on chromosome arm 5DS. The QTLs and the associated molecular markers identified in this research can be used to develop wheat cultivars with potentially durable leaf rust resistance.     

Chromosomal location of genes controlling short-term and long-term somatic embryogenesis in wheat revealed by immature embryo culture of aneuploid lines

The expression of essential genes during somatic embryogenesis can be analysed by inducing aneuploid cells to undergo embryogenesis during immature embryo culture and then determining whether defects occur. Triticum aestivum disomic and aneuploid stocks, including 36 ditelosomics and 7 nullitetrasomic Chinese Spring wheats, were compared for their ability to undergo somatic embryogenesis after 2 months of in vitro immature embryo culture. Their regeneration capacity was observed after 4 and 14 months of in vitro culture to determine which chromosome arms influence the process. The large range of variation found among the tested aneuploids suggested that genetic control of the somatic tissue culture ability is polygenic. Our results indicate that genes affecting somatic embryo-genesis and regeneration are located in all of the homoeologous chromosome groups. The lack of chromosome arms 1AL (DT 1AS) and 3DL (DT 3DS) practically suppresses somatic embryogenesis, demonstrating that major genes on wheat chromosome arms 1AL and 3DL control regeneration capacity. Results suggest that plants were mainly produced from somatic embryo development. Although the control of somatic embryogenesis and regeneration is polygenic, the genes located on the long arms of homoeologous group 3 chromosomes have a major effect. We also have evidence of chromosome arms that determine the time required for regeneration.     

Mapping of glutenin and gliadin genes located on chromosome 1B of common wheat

Summary The inheritance of the high molecular weight (HMW) glutenins and of several gliadins controlled, respectively, by the long and short arms of chromosome 1B of common wheat was studied. Analysis was carried out on the progeny of two inter-varietal crosses in which the parental lines possessed differentially migrating subunits as revealed by sodium dodecyl sulphate polyacrylamide gel electrophoresis. No recombination event was detected either within the fraction of the HMW glutenins or among most of the gliadin subunits studied indicating that they are controlled by tightly linked gene clusters. One gliadin subunit (B30) showed 25.5% recombination frequency with the rest of the gliadin subunits and 23.5% recombination frequency with the fraction of the HMW glutenin subunits. It has been concluded that this subunit is controlled by a separate locus (Gld-B6), proximal to the major gliadin gene cluster on the short arm of chromosome 1B. Consequently, the recombination percentage between the glutenin loci and most of the gliadin loci was calculated as 49.0 and the distance in centi-Morgans (cM) as 53.6. The estimated distance in cM is very close to the observed recombination percentage. A genetic map of these storage protein genes is presented.     

Molecular mapping of genomic regions associated with wheat seedling growth under osmotic stress

A quantitative trait loci (QTL) approach was applied to dissect the genetic control of the common wheat seedling response to osmotic stress. A set of 114 recombinant inbred lines was subjected to osmotic stress from the onset of germination to the 8^th day of seedling development, induced by the presence of 12 % polyethylene glycol. Root, coleoptile and shoot length, and root/shoot length ratio were compared under stress and control conditions. In all, 35 QTL mapping to ten chromosomes, were identified. Sixteen QTL were detected in controls, 17 under stressed conditions, and two tolerance index QTL were determined. The majority of the QTL were not stress-specific. In regions on five chromosome arms (1AS, 1BL, 2DS, 5BL and 6BL) the QTL identified under stress co-mapped with QTL affecting the same trait in controls, and these were classified as seedling vigour QTL, in addition to those expressed in controls. Tolerance-related QTL were detected on four chromosome arms. A broad region on chromosome 1AL, including five QTL, with a major impact of the gene Glu-A1 (LOD 3.93) and marker locus Xksuh9d (LOD 2.91), positively affected root length under stress and tolerance index for root length, respectively. A major QTL (LOD 3.60), associated with marker locus Xcdo456a (distal part of chromosome arm 2BS) determined a tolerance index for shoot length. Three minor QTL (LOD < 3.0) for root length and root/shoot length ratio under osmotic stress were identified in the distal parts of chromosome arms 6DL (marker locus Xksud27a) and 7DL (marker locus Xksue3b). Selecting for the favourable alleles at marker loci associated with the detected QTL for growth traits may represent an efficient approach to enhance the plants’ ability to maintain the growth of roots, coleoptile and shoots in drought-prone soils at the critical early developmental stages.     

Comparison of homoeologous group-6 short arm physical maps of wheat and barley reveals a similar distribution of recombinogenic and gene-rich regions

Eighty two new loci, mapped with 51 DNA clones, were added to the earlier deletion maps of the homoeologous group-6 short arms of hexaploid wheat ( Triticum aestivum L. em Thell., 2n = 6 x = 42, AABBDD). There are now 41, 56 and 52 loci mapped on deletion maps of 6AS, 6BS and 6DS, respectively. The linear order of orthologous loci in all three arms appears to be identical. The majority of the loci are located in the distal one-half of the three arms. There seems to be an increased marker/gene density from the centromeric to the telomeric regions in each arm, and the marker density in comparable physical regions is similar on all three maps. Recombination is not uniformly distributed along the chromosome arms; 60% of recombination occurs in the distal one-third of each arm. Recombination increases from the proximal region to the distal end in a nonlinear pattern. The distribution of loci and recombination along each of the three chromosome arms is highly correlated. Comparison of the 6BS deletion map from this study and a 6HS physical map of barley ( Hordeum vulgare L., 2n = 2 x = 14, HH) reveals a remarkably similar distribution of recombinogenic and gene-rich regions between the two chromosome arms, suggesting that the distribution patterns of genes may be conserved in the homoeologous group-6 chromosome short arms of wheat and barley. A consensus map of wheat group-6 short arms containing 46 orthologous loci was constructed. Comparison of the consensus map with published linkage maps of Triticeae group-6 chromosome short arms indicates that the linear order of the loci on the maps has been largely conserved. Evidence from this study does not support the existence of a 2BS-6BS reciprocal terminal translocation.